Variations in total brain mass and in the mass of three brain regions (main olfactory bulb, hippocampus, auditory nuclei) were examined using a data set for 63 species of bats (Chiroptera). Using both conventional and phylogenetically based analysis of covariance (log body mass as covariate), we tested several hypotheses that relate total brain mass or the size of the components to variation in foraging ecology, categorized as phytophagous, gleaner, and aerial insectivore. In some analyses, the category phytophagous was split into phytophagous pteropodid and phytophagous phyllostomid to examine differences between two distinct clades of bats. Because the Megachiroptera orient primarily by vision and olfaction, whereas all other bats rely on laryngeal echolocation to locate their prey, we hypothesized that the former would differ in size of the main olfactory bulb, as compared with all other bats. This hypothesis was supported by our analyses. Our more general prediction was that insectivorous bats, which rely heavily on echolocation for the pursuit and capture of their prey, would have larger auditory nuclei than do phytophagous species. This, too, was supported. We also compared phytophagous (fruit or nectar consuming) bats in two families, the Pteropodidae and the Phyllostomidae. We hypothesized that the phyllostomids, which use echolocation while foraging, would have larger auditory nuclei. Although statistical power is low in phylogenetically informed comparisons of the two clades, we did find weak evidence in support of this hypothesis. We conclude that bat brains show evidence of adaptation to foraging ecology.

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Using single-copy DNA hybridization, we carried out a whole genome study of 16 bats (from ten families) and five outgroups (two primates and one each dermopteran, scandentian, and marsupial). Three of the bat species represented as many families of Rhinolophoidea, and these always associated with the two representatives of Pteropodidae. All other microchiropterans, however, formed a monophyletic unit displaying interrelationships largely in accord with current opinion. Thus noctilionoids comprised one clade, while vespertilionids, emballonurids, and molossids comprised three others, successively more closely related in that sequence. The unexpected position of rhinolophoids may be due either to the high AT bias they share with pteropodids, or it may be phylogenetically authentic. Reanalysis of the data with varying combinations of the five outgroups does not indicate a rooting problem, and the inclusion of many bat lineages divided at varying levels similarly discounts long branch attraction as an explanation for the pteropodid-rhinolophoid association. If rhinolophoids are indeed specially related to pteropodids, many synapomorphies of Microchiroptera are called into question, not least the unitary evolution of echolocation (although this feature may simply have been lost in pteropodids). Further, a rhinolophoid-pteropodid relationship--if true--has serious implications for the classification of bats. Finally, among the outgroups, an apparent sister-group relation of Dermoptera and Primates suggests that flying lemurs do not represent the ancestors of some or all bats; yet, insofar as gliding of the type implemented in dermopterans is an appropriate model for the evolution of powered mammalian flying, the position of Cynocephalus in our tree indirectly strengthens the argument that true flight could have evolved more than once among bats.

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